The Allopatric Model and Phylogeny in Paleozoic Invertebrates Author(s): Niles Eldredge Reviewed work(s): Source: Evolution, Vol.

25, No. 1 (Mar., 1971), pp. 156-167 Published by: Society for the Study of Evolution Stable URL: http://www.jstor.org/stable/2406508 . Accessed: 09/11/2011 14:15
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THE ALLOPATRIC MODEL AND PHYLOGENY IN PALEOZOIC INVERTEBRATES

NILES ELDREDGE

The AmericanMuseum of Natural History,CentralPark West at 79th,New York, New York 10024 Received June 10, 1970

Time is the one elementof paleonto- cannot be denied that a gradual,strictly change in a specieslogical data whichmitigates, a degree, phyletic, to progressive of the disadvantages inherent the fossiliza- stockhas eventually to populations in led phenotypdistinct tion process. Additionof this fourth di- individualssufficiently mension to evolutionary biology has ically (and probably genotypically)to greatlysharpened out perspective both warrant of of recognition a "new" species,a ratesand modesof evolutionary processes. model which would allow single or mulPaleontologists have understandably em- tiple splittings the lineage into new of phasized the importance time in the species duringthe same time span is the of elaborationof evolutionary models, and more satisfactory the explanationof for have made particularly important contri- the diversity life since the Cambrian. of butionsin the generalarea of the origin On probabalistic groundsalone, we must majority of higher of taxa. The concept gradualism, concludethat the overwhelming an important aspect of geological thinking of metazoan species that have appeared some (see Simpson,1970), has permeated pale- on the earth'ssurfacearose through ontologicthoughtto the extent that all processof splitting. phylogenetic changeis generally conceived analOn the otherhand,paleontological to occur by small increments over vast yses of lineage splittinghave generally periodsof time. This dominantly phyletic dealt only with morphological divergence modelof transformation, stressing im- and do not fullycorrespond neontologthe to portanceof time and the aggregation of ical discussions speciation.A biological of large numbers small steps of morpho- speciesconcepthas been incorporated of into logical change,has underlainmost pale- paleontology recently(see only relatively ontologicaldiscussionsof the origin of generaldiscussions Newell, 1956; Imby new taxa, including species. In fact,this recent studies brie,1957,and twoexcellent phyleticmodel applies not only to strict by Waller,1969; Gould, 1969). Since,in transformation lincases of phyletic (i.e., ear trends in which arbitrarysegments favorable circumstances,paleontologists true are viewed as "new" taxa), but also to have been successfulin recognizing are as valid which of wherenew "biospecies"on criteria mostdiscussions divergence, as tree are con- and complete thoseused to differentiate branchesin a phylogenetic stocksfrom the majorityof recent species, a reapsideredas gradually diverging modelsof specipraisal of paleontological a parentgroup. At the specieslevel,the only such level ation is called for. proin the taxonomic wherea taxon Of the variousmodelsof speciation hierarchy can actually be said to exist in nature, posed and discussedover the past forty such a gradualistic view of the originof years, the allopatric model has gained evnew taxa is, in one sense at least, clearly nearlytotal acceptanceamongcurrent that at odds with currently biologists.I wouldsuggest accepted views of olutionary 'speciationderived from studies of the the allopatricmodel (geographicspeciain recent biota. On the one hand, while it tion) be substituted the mindsof paleEVOLUTION

25:156-167. March 1971

156

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FIG. 1. Outlinemap of a portionof easternand central North America showing the geographic relationshipbetween the epeiric sea (stippled), the marginal shelf sea (cross-hatched), and the land areas (clear). Land bordering bordering the craton to the north was apparentlyeroded to a low level, while the land area to the east was being actively uplifted,perhaps as discontinuousislands in an arc system.

ontologistsfor phyletictransformism as ENVIRONMENTS OF PALEOZOIC EPEIRIC the dominant mechanism the originof of AND MARGINAL CRATONAL SEAS new speciesin the fossilrecord, and that Sedimentsdeposited in the extensive the allopatricmodel,ratherthan gradual Paleozoic seas that coveredmuch of the morphological divergence, the morecor- continental is interior NorthAmerica of have rectviewof theprocesses underlying cases yieldedvast quantities invertebrate of fosof splittingalready documented by nu- sils. These extensiveepeiric seas have merousworkers. been conceived as predominantly of warm, Recognition ancestral-descendant shallowbodies of water in of rewhichgeneral in lationships the fossil recordare often environmental conditions were ratherhobased essentially biostratigraphic on data. mogeneous within broad bands parallel to in Actually,such relationships fact conshorelines (Shaw, 1964); diurnalfluctuastituteuntestable hypotheses.The ancestionsin physicalenvironmental parameters tral-descendent berelationships presented werepresumably negligible whencompared low for the various taxa of the Phacops of rana complexshould be viewed as such to environments the marginalbasins. (basins) and highsprohypotheses. The theory (cladistic) re- Local depressions of to lationships, upon which these hypotheses vided some geographicheterogeneity of ancestral-descendant are these epeiric seas. The general environrelationships based, is set forthin detail in Eldredge mentsrepresented epeiricseas have yet by (1969). to be analyzed fully. Marginal shelfen-

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vironmentsare preserved in generally this discontinuity have been exhaustively thickersedimentary sequences deposited discussed in recent years (for example, in linear troughs, geosynclines, or either interalia, see Newell,1956, 1959; Imbrie, on, or immediately adjacent to, the conti- 1957; Durham, 1967). One particularly nentalmargins. lineages aspectof discontinuous interesting In the Middle Devonian,the sourceof occurs as the general rule in the fossil the inhabiting epeiric the examplediscussedbelow, true epeiric recordof organisms inundations the craton (stable conti- seas. In this instancea given species A, of nental interior) occurred several times whichmay be quite abundantand known area, is confined (see Fig. 1); the primary types of sedi- over a large geographic ments depositedin these seas were car- to one or more bodies of rock strata bonates(limestones) and calcareous shales. throughout geographic range. A very its The MichiganBasin was thechiefcratonal similar closelyrelated speciesB, obviously structure providingfaunal heterogeneityto species A and quite likely descended in the Middle Devonian; for instance, fromit, is foundin all or some of the Imbrie (1959) lists approximately80 areas stratigraphically above species A. the speciesand subspecies articulate of brachi- There are no known intermediates; opods endemicto that area in the Middle discontinuities may not, but often do, Devonian Traverse Group. The epiconti- occur at lithologic boundaries. The supnentalbasin on the easternmargin the porting of of for evidence a hypothesis ancescontinent (Fig. 1), corresponding roughly tral-descendant betweensperelationship to the Allegheny and foldedAppalachian cies A and B is oftenstrengthened the by geomorphologic provinces, preserves a absenceof any otherspeciesof that genus thickersequence of sediments.The an- livingduringthat particulartime period cient shorelineran approximately north- in that particularfaunal province. An south; near-shore sedimentswere domi- example, from literathe drawnat random nantly non-calcareoussilts and sands ture,whichfitsthis generalcharacterizaderivedfrom uplifted the mountains the tion, would be the relationship to between east. Further off-shore(to the west), the Ordovician lorettrilobite Cryptolithus sedimentsbecome finer,and calcareous tensis Whittington and its possible deshales and occasional limestonesappear scendantC. bellulus (Ulrich) (Whittingin the stratigraphic column. Thus, the ton, 1968). Many such examples are familiar with axis of greatestsubstratevariationruns knownto all paleontologists roughlyeast-west,normal to the north- Paleozoic epicontinental sediments. far and extending into the southshoreline basic models the Figure2 illustrates four continental interior.The axis of greatest whichmay be invokedto explainsequenancesof biological variationin organismsstudied tial occurrences non-intergrading vari- tral-descendant species. Variationson all so farparallelsthisaxis of substrate and ability. A more complete discussionof fourof thesemodelsare conceivable, to intended be exhausand envi- the diagramis not Middle Devonian stratigraphy three sucvariation tive. In all fourillustrations, and biotic geographic ronmental lithology cessiverock unitsof unspecified is givenin Eldredge(1969). and are and thickness numbered, the three THE FOSSIL RECORD IN PALEOZOIC speciesare indicatedA, B, and C. Width EPEIRIC SEDIMENTS an of the diagramrepresents axis of morof phological Figure2A showsthe difference. Though nearlycontinuous sequences has which largely evolving lineages are known from the classicmodelof saltation, fossil record,discontinuity by far the falleninto disfavoramong zoologistsand is If morecommonstate of affairs. Causes of paleontologists. the putativesaltations

ALLOPATRIC MODEL

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Fm.3 saltation

Sp. C

Fm.3
migration
.

Sp.C

Fm.2 saltation

sp. B

Fm.2

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MORPHOLOGIC CHANGE

MORPHOLOGIC CHANGE

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MORPHOLOGIC CHANGE

FIG. 2. Four models of speciation used to explain the occurrence threespecies (A-C) in three of successive bodies of strata (Fms. 1-3) in one local area. Width of the diagram is an axis of morphologic change. A. Saltation model. B. Phyletic model. C. Gradual morphological model. divergence D. Allopatricmodel where morphologicalchange occurs relativelyrapidly in peripheralisolates. For fullerexplanation,see text. Solid lines: species occurs in local rock unit; Dashed lines: species is living in the same faunal province but outside the local area. AS: allopatricspeciation. Hiatus indicated by stippling.

take place at or near a formation boundary, the possibilityof a lengthyhiatus is at least tacitlyconsidered be small. to Another implicit consequence thismodel, of

of greater importance here,is the assumptionthatlittleor no morphological change occurs withina species withinits stratigraphicinterval occurrence. of

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models are rarely,if by the nature of most fossiloccurrences While saltationist ever, invokedby modernpaleontologists,in epeiricsediments.As shownin Figure speciesA, B, or C are found model (Fig. 2B) is 2B, wherever the standardphyletic a cited throughout rock sequence,theyare exthe probably processmostcommonly linear (see Newell, pected to show a gradual,generally to explain such occurrences 1956, p. 67ff; Imbrie, 1957, p. 129ff; changein some aspects of theirmorpholex- ogy. It has been claimed (e.g., Imbrie, Durham, 1967, p. 560). An excellent acceptedby ample of analysis along these lines is 1957, p. 143), and implicitly that the relative most paleontologists, (1951, p. 95ff) reinSylvester-Bradley's examplesof such of terpretation Carruthers'(1910) work scarcityof documented through delanouei change within a species-lineage on the rugosecoral Zaphrentites and Haime). Linear se- a sequence of rocks is due largelyto a (Milne-Edwards reasonof lectionpressuresare thoughtto produce scarcity rockunitsrepresenting over a sufsedimentation a gradual (often statistical)change in a ably continuous long period of time. On the characteror suite of characters. Strati- ficiently of the graphicbreaks in the occurrence the contrary, usual case, at least in Paleois of lineageare theresult local establishment zoic epeiricsediments, for the observer the conditions, to documentno change throughout of unfavorable environmental rangeof speciesA, B, or C, as possiblythoughnot necessarily severe stratigraphic model, Fig. 2A). absence of marineconditions, (as in the saltationist as complete in Figure 2B. Non-overlapof Such changesas may be apparentusually the case otherthan the speciescharactertraitsis explainedby invoking involvecharacters eiused to distinguish of continuation the gradual linear trend specificcharacters from in anothergeographic area where condi- therof a pair of lineagecomponents tions remainfavorable. Reappearanceof the other. Perhaps instead of relaxed it of the next segment the lineage recogniz- selectionpressures, would be more acin able in thelocal rockcolumn, thisview, curate to attributethis relativelack of selection. back into changeto stabilizing of is a simplematter migration have seldomemphasized of Paleontologists the local area upon the reinitiation In stability.MacGillavry conditions. the this morphological environmental favorable epeiricseas, thismodelmay well correlate (1968, p. 70) has writtenthat "many Moore species do not show any evolutionary and withtransgressions regressions. (1954, 1955), in fact, claimed that re- change at all." MacGillavrypoints out of gression the epeiricseas onto the mar- that, althoughgradual change mightbe basis, it is rarely on shelvescreatedcrowded expected a theoretical ginal continental and selectionpressures, encountered, that such a situationis intensified conditions, process. of and resulted in higher (morphological) not an artifact the fossilization to seen during Anotherof the rare statements this rates than were evolutionary The effectwas made by Kurten (1965, p. thetimesof widespread transgression. basic phyletic model, however, plainly 345): "The situationsuggeststhat new rapidly,but underliesthe specificmodel that Moore species arose comparatively withtendedto continue once established, proposes. Though it will be argued below that out any change." Actual documentation the origin of many, if not most, new for this alleged stabilizingselectionand lack of linear morphological species in the Paleozoic occurredwithout concomitant of and to reference transgressions regressions, change is as rare as documentation Moore's claim of relaxed evolutionary such change,thougha similarobservation inun- seems to underlieMoore's (1954, 1955) rates duringperiods of widespread dationof thecratonseemsto be supported arguments.

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Stenzel (1949) applied a gradualistic, descendants, keepingwith the general in linearmodelto explainthe non-intergradsituation shown in Figures hypothetical ing sequentialsuccessionof four species 2A and 2B. Many examples,of course, of the Middle Eocene oysterCubitostrea. are knownwhere descendantspecies are Depositional hiatuses separate the four foundin the same strata,possiblysymspecies in this succession,and though patric, species. ancestral withthepresumed trends are apparent throughthe entire divergence,as Gradual morphological lineage,the fourspeciesdo not intergrade. discussedabove, is not in any respecta Though not explicitly stated by Stenzel, strictalternative allopatric,sympatric, to it is apparentfrom discussion his that no or stasipatric (White, 1968) models of progressive trendsin species-specific char- speciation. Formulation of discussions acters were documentedon the intra- (and illustrations branching tree diaby specificlevel withinthe stratigraphic in- grams) of lineage-splitting terms of in terval of occurrence any of the four gradual morphological of divergencesimply species. As interpreted here, then,Sten- reflects predithe typicalpaleontological zel's "successionalspeciation"in the Cu- lectionfor the phyleticmodel of gradual bitostreastock does not seem to fit the change (Figure 2C). The assumption of model of gradual phyleticevolutionwith gradual morphological divergenceis so gaps in the record(Fig. 2B). that thinking pervasivein paleontological A somewhat similarcase was presented lineage-splitting oftenbecomesreducedto by Kaufmann (1933) in his belatedly a special case of the phyletic model. celebratedstudy of the Upper Cambrian Rarely does the paleontologist consider trilobite Olenus. Kaufmann recognized how the splittingis actually effected; that,whilesome characters changedgrad- though as paleontologists a wholeare not ually within speciesand between a species ignorantof the allopatric model, it is of a singlelineage,othersremainedcon- nevertheless true that many discussions stantwithin of species,and changedabruptly up to the presentday presentsplitting fromspecies to species. Another example a lineage as if it could have occurred of stability (sub-)-species-specific in char- gradually timewentby within single a as actersin marine of invertebrates theepeiric local area of investigation.In recent seas is presentedbelow. Further docu- discussions(e.g., Kermack,1954; Imbrie, in mentation support thishypothesis of 1957) of the splitting the Upper Cretaof of morphological stability is desirable. If ceous Micraster Wright stock,M. corbovis true,suchstability automatically precludes and M. senonensis Lambertare regarded the use of the phyleticmodel of species as gradual offshoots fromthe main Miat transformation, least in its simplest craster phylum;whileKermack (1954, p. in of form, the analysisof the origin most 422) hints at establishment reproducof such invertebrate species. tive barriersbetweeneach of these two and the main line in the north Generallyspeaking,only in such in- offshoots stanceswherean ancestral dispeciesis found of England, an essentially sympatric in strataalong with,or at least the same vergence at least implied thesouthern in is age as, a presumeddescendantspecies, area. have paleontologists discussedsplitting of Referencesto "geographicspeciation" a species-lineage. Figures2C and 2D illus- or "geographic are subspeciation" not untrate persistence ancestralspecies into commonin the recentpaleontological of litthe time range of the descendants; in erature. Figure 2C can easily accommoboth of thesediagrams, ancestralspe- date gradualmorphological the with divergence cies are depictedas not livingin the same the allopatric spemodel,withdescendant area (i.e., the local rock column) as the cies simplymigrating back into the local

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pending area of interest. But the retentionof these questionsare best deferred exampleof of gradual divergence, emphasizing time a discussion a documented the of element, remains odds withthe general allopatricsplitting a Paleozoic speciesat pictureof morphological stability Pale- lineage. of ozoic invertebrates the epeiric seas. of THE PhacopsranaLINEAGE OF THE Figure 2D depictsthe most likelymodel MIDDLE DEVONIAN for this general situation. In terms of the species-specific morphological features Two speciesof the trilobite genusPhawhichserveto distinguish two species cops Emmrichare foundin the Middle any of an ancestral-descendant lineage, most Devonian GivetianSeries of easternand evolutionary change occurs allopatrically. centralNorthAmerica(see Figures3, 4). While linear selection (gradual morpho- Analysisof a large body of morphologic, logicalchange) may occurwithin any one stratigraphic, geographic data has led and species throughtime, such change does to the conclusion that both Phacops rana not necessarily, usually does not, in- (Green,1832) and Phacops iowensis and Delo, volve species-specific morphological fea- 1935 were true "bio-species" (Eldredge, turesin whichthe two speciesdiffer.An 1969). Of the two,P. iowensis was almost important exception this generalization certainly to a from phacopidstock descended occurswhena descendant speciesbecomes presentin North Americaat least from sympatric with its ancestralspecies. In Lower Devonian Gedinniantimes. Presthis case, numerous interactions, including ence of the stock in other parts of the hybridization characterdisplacement, worldhas yet to be established.On the and can occur which have important effects otherhand,P. rana is mostcloselyrelated on such morphological features.All evo- to P. schlotheimi (Bronn) of the Eifelian lutionary phenomena knownto affectthe Series of West Germany, and a closely morphology individuals of within species related form from the Spanish Sahara a (mimicry, characterdisplacement, clines, (Eldredge, 1969; C. J. Burton, pers. etc.) are potentially documentable the comm.). P. rana and P. iowensishave in fossilrecord (Eldredge, 1968, 1969; Van about the same time-stratigraphic range Valen, 1969). Generallack of recognition (Givetian-representing 8 approximately of these phenomenato date is one con- to 10 million years). The interactions comitant the failureto incorporate of the betweenthese two species were complex allopatricmodel fullyinto paleontological and are discussed in detail elsewhere thinking. (Eldredge,1969). In termsof the specificexample of a While phacopids on the whole were discontinuous as species-lineage typically morphologically of a uniform, number anain preserved Paleozoic epeiricmarinesed- tomical featuresshow consistentdiffera iments, purelyallopatricmodel (Figure ences between the two North American the 2D) is best suitedto explaining origin Givetianspecies. Of these,the most imof the descendant taxa. Two further ques- portant or involvethe schizochroal, aggretions remain: can we identifygeneral gate, eye. Such eyes are ratherunique in (and arthropods genareas where marginalpopula- among trilobites geographic tionsmay becomeisolated,and new sub- eral) in that each lens is coveredby a speciesand speciesappear? What,if any, separate cornea. The lenses are aggrein is the relationship betweenallopatrically- gated on a visual surfaceand arranged and the many verticalcolumns,or dorso-ventral (d.-v.) based evolutionary changes of and transgressions regressions theepeiric files, (Clarkson,1966). A stable number each files,characterizing seas that occurredduringthe time spans of dorso-ventral withina phacopid speof many species lineages? Answers to population-sample

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EUROPE FRASNIAN

NORTH AMERICA SENECAN TAGHANIC

FORM TULLY

TIOUGHNIOGA GIVETIAN ERIAN

MOSCOW
LUDLOWVILLE SKANEATELES

... cephalon ...B..... 1.5. Letltra.iw showing.. lese arranged. in..dorso-ventra... 2. ..... ....... ... 3...I...

ofcpao

files,

CAZENOVIA MARCELLUS

cies. reahein.. is. . ... early holspi ontogeny. mainder ontogeny. Dorso-ventral of Z. ..... file~~~~~~~~~~~~~~~~~~~~~~~~~~~~2'Z= FIG..4. Comparison of time-stratigraphic nonumberthesingle is imotntapc ~~~most; menclaturesfor the Middle Devonian in Europe .. ofbothinerpoultina and.inter.specifi ......... and North America, and the rock-stratigraphic variation the.Phcopida within Most nomenclature the "type" section of the North of ofth. lihtmrpoogca.hag AmericanMiddle Devonian. wit .ownsi.thouhou.it.stat seen.... P... grphc.ane.asaparnty.hyetc. ..n. insar.otrs,.hw.amc grete. aout f.voutonrychng changesare best given subspecific status. withi sam the tmespn.om.o.wic is.ls apparently ttributableto.gradual The discussion,then, will centeraround phyletic ..evouton.Mstof.heevluio the pattern allopatrically-based of changes witintheP.raa.soc,.oweer.i ... volved chage in.orpolgicl.eatre in characterstates. The characterstate in populations near the periphery of the~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~. ....... geographic. rag.ftespce n r whichmostclearly undergoes allopatrically thuallopatric.in nature..Theapparent based changeis the number dorso-venof phyletic. chngswihn.h.sok.o tral files. ever,indicate that total geneticisolation~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~... ... probably.. did.no.occur.and.fr.thisreaso file Reductionof dorso-ventral number the. taxareconizd.onthe.asi new... from18 to 15 was the major evolutionary of ..the. a.optrcalybse.mrpolgia change in the P. rana stock throughout its history.The basic evolutionary pattern of this reduction involvedgeographically which isolatedtransitional populations gave rise to populations havinga new,reduced files. Populations number dorso-ventral of isolated remained withthisnew filenumber the fora timefrom populations exhibiting thengradually ancestral condition, migrated over the species range and replaced the moreprimitive populations.A longperiod reof stabilitythen ensued until further and the more duction was accomplished primitive populations wereagain replaced. The oldest examplesof P. rana known in occuras rarespecimens theLowerCazenovian Stage Dundee and Delaware limeMember stonesof Ohio,and in theSolsville in of theMarcellusFormation central New York. They are uniformly characterized

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by 18 dorso-ventral files,a featurethey in age. All Phacops rana fromthis forhave 17 d.-v.filesand are attributshare with their putative European an- mation cestor, P. schlotheimi.P. rana popula- able to P. rana rana, and all 17 d.-v. file tions with 18 d.-v. files persistinto the populationswere limited to the eastern the Upper CazenovianSkaneateles equivalents marginalsea throughout durationof generally sea westof the marginal of New the CazenovianStage. The widespread basal Tioughniogan York and the Appalachians. They occur in Skaneateles equivalents in Michigan Centerfield faunamarksthe disappearance (Bell shale, RockportQuarry limestone, of all 18 d.-v. file P. rana and the first of Ferron Point Formation,Genshaw For- incursion 17 d.-v. file P. rana west of sea. P. rana has 17 mation), Illinois (St. Laurent limestone), the easternmarginal extent its Indiana (Deputy and Silver Creek lime- d.-v. filesthroughout geographic stones), Ohio (Silica shale), and south- duringthe entireTioughniogan. files A further reduction dorso-ventral in western Ontario(upper and lowerArkona the during TaghanicStage.Again, shale). In addition, theyare knownfrom occurred occurs transitional population float material in Iowa and from lower an evidently sea Hamilton rocks in the southernAppa- in the marginal of the east. Specimens of lachians. Two eye variants, involving pri- fromthe Tully limestone centralNew in marily differences number lensesper York show a range of 15 to 17 dorsoof dorso-ventral have led to recognition ventralfilesin samplesdrawnfrommost file, of two sub-taxa withinthe 18 d.-v. file localitiesin centralNew York. It is the group:P. rana milleri Stewart and P. rana only otherinstancewherethe numberof withassurcrassituberculata Stumm. Numberof lenses d.-v. filescannotbe predicted to one specimen the next. Loss per d.-v. file is a componentof inter- ance from files is evidentlypedopopulational variation not encountered of dorso-ventral in since the largerspecimens the again in the subsequent history P. rana, morphic, of in populations both the Solsand the generalnatureof this geographi- transitional of callybased variability notbe discussed ville and Tully limestones New York will completend to reach the moreprimitive in further this paper. files. Additionof of Studyof the growth theeye of popu- ment of dorso-ventral retarded lationswith18 dorso-ventral suggests the finald.-v.file(s) is seemingly files later that the first(anterior)fileis added last intoprogressively stagesof ontogeny, lost. However, in holaspidontogeny. This fileis reduced until they are ultimately quantum to a singlelensin theone specimen known loss of d.-v.filesdoes not involve fromthe Lower Cazenovian lower shale deletionof a file; populationsexhibiting unit of the SolsvilleMemberof the Mar- variabilityin d.-v. file numbershow a of cellus Formation centralNew York. A continuous gradation from fully defileswitha reduced small sample from the overlyingsand- velopedfiles,through stone unit of the Solsville shows great numberof lenses,to total deletion. is resolution currently Biostratigraphic of variationin the development the first d.-v. file. One specimenshows the file inadequate to documenta time lag bede- tween the appearanceof 15 d.-v. file P. it whereas is partially fullydeveloped, veloped or totallyabsent in others. This rana in the easternmarginalsea and its a spread over the sample seems to represent transitional (presumably)subsequent in interior theepeiricsea. Fifteen phase in the P. rana lineage. The next cratonal the unit younger d.-v. fileP. rana from TaghanicStage succeedingPhacops-bearing to P. rana norwoodensis than the Solsville in New York is the are all referable Stafford whichis eitherupper- Stumm. This subspeciesoccurs in most limestone, known(upper Cedar Skaneateles Taghanic formations most Marcellus or lowermost

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Valleyof Iowa; Milwaukeedolomite, Wisconsin;upperPetoskey, western Michigan; Tully limestone, New York). No P. rana are knownfromthe Taghanic of eastern Michiganor northern Illinois. The one specimen P. rana of possible of post-Givetian age available ("P. nupera" Hall-A.M.N.H. No. 496911) is poorly preservedbut apparentlyhas 17 dorsoventralfiles. This may indicatethat the main stockof P. rana rana survived a for while in the marginalsea in post-Tully times afterhaving given rise to P. rana norwoodensis.

in and intergradation the show variability files of number dorso-ventral seem to repshortspans of time,parresentrelatively ticularlyinsofaras the Solsville population (18-17 d.-v. files) is concerned. trend, Rather than a gradual progressive reductionin file numberseems to have occurredratherrapidly. There was an intervalof fromone to two million years between the time of firstappearance in the marginalsea of 17 d.-v. file P. rana and its spread over the craton in the epeiric seas. During muchof thistime,the Cazenoviansea was over the eastern quite extensive generally PhacopsranaAND THE ALLOPATRIC MODEL one-halfof the North Americancraton Reductionin numberof dorso-ventral (Cooper, 1957). The originof 17 d.-v. filesinvolved origin a newcharacter fileP. rana does not appear to be correthe of state in a certain geographicarea, and lated with a period of widespread reits subsequent spread over the species gression,as might be supposed under range,on two separateoccasions. In the Moore's (1954, 1955) model. However, first instance, expansion in geographic the spread of the new characterstate in Stage, and again rangelaggedwell behindthe origin the the basal Tioughniogan of new state,but the spread of the 15 d.-v. in the Taghanic Stage, seems to coincide file conditionin the Taghanic was "in- with a renewed,large scale episode of new Many faunal elements stantaneous"in terms of our biostrati- transgression. to thecratonalseas made theirappearance graphicresolution. The distribution populations of charac- in both the early Tioughniogan and seas. It wouldseemlikely, terized a givennumber d.-v. filesin Taghanicepeiric of by epeiric sediments, to then, conforms the then,that the appearanceof a new d.-v. resimply hypotheticalsituation discussed earlier filevariantin epeiricsediments flects migrationin conjunctionwith a (Figure 2). Figure 5 summarizes histhe a whichfollowed widetoryof P. rana, and is mostcloselycom- majortransgression parableto Figure2D. Rocksof Cazenovian spread regression(especially well docuage in the midwest produce 18 d.-v. file mentedbelow the Taghanic Stage) that of for P. rana; these rocksare commonly over- accounted thedisappearance populanumberof tions with the more primitive lain by a basal Tioughniogan correlative of the Centerfield of limestone New York, files. No claim is made for extinction between populations and all P. rana known fromsuch units due to competition have 17 dorso-ventral files. No samples bearing the ancestral and the derived that exhibit variation and intermediate characterstates. of history P. rana (see The evolutionary are in conditions d.-v. file number known fromany of the epeiric sectionsof the Fig. 5), then,fits the situationdepicted midwest.There is no progressive change in Figure 2D very closely. Though the the here represent pattern seen in this or any othercharacterstate data presented in throughout any given formation;this is of splitting but a single stock of the that populated true also of samplesof P. rana foundin myriadsof invertebrates in in particularformations the easternmar- the Paleozoic seas, the similarity modes of of occurrence elementsof this stock, where ginalsea. The twoinstances samples

166
EPEIRIC SEA

N. ELDREDGE
MARGINAL SEA

+
TAGHANIC STAGE

TX
-' - ; -,

TIOUGHNIOGA STAGE

++

CAZENGVAI

onous conditions evidently represented by manyof the Paleozoic epeiricseas. If P. rana is any indication, is in the sediit mentary recordof the seas on the continentalmarginsthat we should look for the origin of new taxa, an unfortunate situationin view of the great amountof deformation consequently and poorerfossil recordusually foundin such areas. But, if this view is correct, documentation the of phylogenies the Paleozoic inverteof brate taxa will remain incompleteuntil the significance the allopatric model of is trulyrealizedand the sediments the of marginal seas bordering cratonalareas the receivethe carefulscrutiny they deserve.
SUMMARY

STAGE

Emphasison timeand gradualistic transformation led to the dominance a has of strictlyphyleticmodel of species transFIG. 5. Phylogenyof the Phacops rana stock in thought. in the Middle Devonian of North America.Num- formation mostpaleontological cases of lineage-splitting bers at the base of the diagram referto the pop- Even documented ulation number of dorso-ventral files. Dotted are often interpreted by recourse to a lines: origin of new (reduced) number of d.-v. gradual (morphological) divergence model files in a peripheral isolate; horizontal dashed in which strong a element phyletic of thinklines: migration; vertical solid lines: presence of taxon in indicated area; dashed vertical lines: ing is incorporated. Allopatricspeciation,not a strict alpersistenceof ancestralstock in a portion of the marginal sea other than that in which the de- ternative gradual divergence, to seems to rived taxon occurs. Crosses denote final disap- fitthecommon pattern non-intergrading of pearance. Compare figure2D; for fullerexplanaspecies withina lineage as typically pretion,see text. servedin Paleozoic epeiricsediments. The majorityof species preservedin epeiric (and absenceof any recognizable gradual sedimentsshow no change in speciesphyletic changewithin them), to the gen- specific the characters throughout interval eral patternsobservedin other inverte- of theirstratigraphic and the occurrence, bratesindicates thatit may well represent phyleticmodel is inapplicableto most of a basic patternof evolution record. applicableto theseelements thefossil of Instead, manyPaleozoic invertebrates. the allo- change in, or developmentof, speciesIf patric model is to be applied at all to specificcharactersare envisionedas ocit Paleozoic invertebrates, I think must, curringrelativelyrapidly in peripheral as the lineargeosynclinal the isolates. Morphological belts bordering of stability epeiric craton, representing perhaps more open species is attributed stabilizingselecto marine conditionsthan the epeiric seas, tion. are not a priorithe morelikelyareas for The Devonian trilobitePhacops rana the establishment peripheralisolates. illustrates, two different of at times in its Yet the shallow shelf environment may history, origin a newcharacter the of state well have represented moreheterogeneous in peripheral isolates. Most new Paleozoic than environments the widespread, monot- invertebrate probably taxa arosein geosyn15 17 18 18 17 16 15

ALLOPATRIC MODEL IN PALEOZOIC

INVERTEBRATES

167

clines (marginalseas) bordering crathe tons; though these originationevents themselves were probablynot related to marineregressions, invasionof new taxa into the epeiric seas seems directlyrelated to periodsof transgression.

Olenus Dalm. Abh. Geol.-Pal. lobitengattung Inst. Univ. Greifswald.10:1-54. K. KERMACK, A. 1954. A biometricalstudy of and M. (Isomicraster) Micraster coranguinum senonensis. Phil. Trans. Roy. Soc. London, B. 237:375--428. KURTEN, B. 1965. Evolution in geological time. In Moore, J. A. (ed.), Ideas in Modern Biology, Natural History Press, New York: 329ACKNOWLEDGMENTS 354. H. I thankR. L. Batten,J. C. Boylan,S. MAcGILLAVRY, J. 1968. Modes of evolution Bijdragen mainly among marine invertebrates. J. Gould, G. J. Nelson, N. D. Newell, tot de dierkunde.38:69-74. H. B. Rollins,and J. Spillerforcomments MOORE,R. C. 1954. Evolution of late Paleoon an earlierversionof the manuscript, zoic invertebrates response to major oscilin and G. R. Adlington the photographs. lations of shallowseas. Bull. Mus. Comp. Zool. for 112:259-286. . 1955. Expansion and contractionof shalLITERATURE CITED low seas as a causal factorin evolution. EvoR. G. 1910. On the evolution of CARRUTHERS, lution 9:482-483. Zaphrentis delanouei in Lower Carboniferous NEWELL, N. D. 1956. Fossil populations. In times. Quart. J. Geol. Soc. 66:523-538. P. Sylvester-Bradley, C. (ed.), The Species E. N. K. 1966. Schizochroal eyes CLARKSON, Concept in Palaeontology. SystematicsAssoc. and vision of some Silurian acastid trilobites. Publ. 2:63-82. Palaeontology9:1-29. . 1959. Adequacy of the fossil record. J. COOPER, G. A. 1957. Paleoecology of Middle Paleontology33:488-499. Devonian of easternand centralUnited States. SHAW,A. B. 1964. Time in Stratigraphy.McGeol. Soc. Amer.,Mem. 67:249-278. Graw-Hill,New York. DURHAM, J. W. 1967. The incompletenessof SIMPSON, G. G. 1970. Uniformitarianism. An our knowledge of the fossil record. J. Paleinquiry into principle,theory,and method in ontology41:559-565. geohistoryand biohistory. In Hecht, M. K. ELDREDGE, N. 1968. Convergence between two and W. C. Steere, (eds.), Essays in Evolution Pennsylvanian gastropod species: a multiand Genetics in Honor of Theodosius Dobzvariate mathematicalapproach. J. PaleontolNew York: hansky. Appleton-Century-Crofts, ogy 42:186-196. 43-96. 1969. Geographicvariation and evolution STENZEL, B. 1949. Successionalspeciation in H. in Phacops rana (Green, 1832) and Phacops paleontology: the case of the oysters of the iowensis Delo, 1935, in the Middle Devonian stock. Evolution 3:34-50. sellaeformis of North America. Ph.D. thesis, Columbia SYLVESTER-BRADLEY, P. C. 1951. The subspeUniversity. cies in palaeontology. Geol. Mag. 88:88-102. S. J. 1969. An evolutionary micro- VAN VALEN,L. 1969. Variation geneticsof exGOULD, cosm: Pleistocene and Recent history of the tinct animals. Amer. Natur. 103:193-224. land snail P. (Poecilozonites) in Bermuda. WALLER,T. R. 1969. The evolution of the Bull. Mus. Comp. Zool. 138:407-532. Argopecten gibbus Stock (Mollusca: BivalIMBRIE, J. 1957. The species problemwith fosvia), with emphasis on the Tertiary and sil animals. In Mayr, E. (ed.), The Species Quaternaryspecies of eastern North America. Problem. Amer. Assoc. Adv. Sci. Publ. 50: Paleont. Soc. Mem. 3 (J. Paleontology,43(5) 125-153. supp.): 1-125. . 1959. Brachipods of the Traverse Group WHITE, M. J. D. 1968. Models of speciation. (Devonian) of Michigan. Bull. Amer. Mus. Science 159:10,65-1070. Nat. Hist. 116:351-409. H. B. 1968. Cryptolithus(TriWHITTINGTON, Unlobita): specificcharactersand occurrencein KAUFFMAN, R. 1933. Variationsstatistische the Ordovician of eastern North America. J. tersuchungeniiber die "Artabwandlung"und Trian Paleontology42:702-714. "Artumbildung" der oberkambrischen